The Importance of Independent Data

Sets

Multiple data sets

support the idea
that species have
descended, with
modification, from
a common
ancestor

Internal Consistency
Internal consistency is
The observation that data from independent sources
agree in supporting predictions made by a theory
Perhaps the most powerful evidence for any scientific
theory, including evolution by natural selection

Internal Consistency in
Cetacean Evolution
Evidence for the evolution of the
cetaceanswhales and dolphins
from terrestrial ancestors
illustrates the idea of internal
consistency
Fossil cetaceans have unique
ear bones
A phylogeny of cetaceans
Relative and absolute dating
DNA analysis
Comparison of vestigial hip
and limb bones in adult
whales and dolphin embryos

Internal Consistency
Data from many different sources are much more consistent
with evolution than with special creation
Descent with modification is a more successful and powerful
scientific theory
It explains observationssuch as vestigial traits and the
close relationships among species on neighboring islands

How Does Natural Selection Work?

Many researchers had proposed the fact of
evolution,
Theory Evolution
Pattern Descent with modification
Process - ?
Natural Selection
Darwins contribution was describing a process,
natural selection

Darwins Inspiration
Darwin based his idea of natural selection on
A model system pigeon breeding
and
The work of Thomas Robert Malthus

Darwins Inspiration
Artificial selection in pigeon breeding
Darwin crossbred pigeons and observed how
characteristics were passed on to offspring
He concluded that the diverse pigeon breeds all
descended from wild pigeons

Darwins Inspiration
Thomas Robert Malthus wrote the book An Essay on the
Principle of Population
Described a struggle for existence
Many more individuals are born than can survive
Thus, people compete for resources

Darwins Inspiration
Darwin combined several ideas to arrive at his concept of
natural selection
His observations of artificial selection
The notion of struggle for existence in natural populations
Variation in natural populations

Darwins Inspiration
Alfred Russel Wallace (1823-1913)
Naturalist
Explorer
Geographer
Anthropologist
From his studies in the East Indies, he proposed the same
ideas as Darwin.
Darwin and Wallace both published their findings in the
Proceedings Of The Linnaean Society of London .
Both Darwin and Wallace arrived at the same idea, but
Darwins name is widely associated with the concept of
natural selection

Darwins Four Postulates

Darwin broke the process of evolution by natural
selection into four criteria, or postulates
1. Individuals in a population vary in their traits
2. Some of these differences are heritable; they are
passed on to offspring
3. In each generation, many more offspring are
produced than can survive
4. Individuals with certain heritable traits are more likely
to survive and reproduce

Darwins Four Postulates

The selected traits will increase in frequency in the population
from one generation to the next, causing evolution
Evolution is a change in the allele frequencies of a
population over time
Evolution is thus a logical outcome of the four postulates
Modern biologists condense Darwins four steps into two
statements:
Evolution by natural selection occurs when
Heritable variation leads to
Differential reproductive success

Testing Darwins Postulates against the

Tuberculosis, antibiotic resistance example from
lecture 1

Variation existed in the population

Due to mutation, both resistant and nonresistant strains of TB were
present before administration of the drug
The variation was heritable
The variation in the phenotypes of the two strains was due to variation
in their genotypes
There was variation in reproductive success
Only a tiny fraction of M. tuberculosis cells survived the first round of
antibiotics long enough to reproduce
Selection occurred
The cells with the drug-resistant allele had higher reproductive success

Biological Definitions
Biological fitness
The ability of an individual to produce surviving, fertile
offspring relative to that ability in other individuals in
the population
Adaptation
A heritable trait that increases an individuals fitness in a
particular environment relative to individuals lacking that
trait
Selection
Differential reproduction as a result of heritable variation

Lecture 3: Evolutionary Process

Freeman: Chapter 23
Dr. Gurney

Lecture Overview

Lecture Outcomes
Test whether evolution or nonrandom mating is occurring
at a particular gene, using the Hardy-Weinberg principle.
Describe the four evolutionary mechanisms, and list
whether each one causes adaptation, introduces new
alleles, acts randomly, or causes genetic variation to increase or decrease.
Explain how nonrandom mating can affect genotype
frequencies without causing evolution.
Define sexual selection. Explain which sex is more
strongly affected by sexual selection, and why.

Introduction
There are two fundamental messages in this chapter:
1. Natural selection is not the only agent responsible for
evolution
2. Each of the four evolutionary processes has different
consequences

Introduction
There are four mechanisms that shift allele frequencies in
populations:
1. Natural selection increases the frequency of those alleles
that contribute to reproductive success in a particular
environment
2. Genetic drift causes allele frequencies to change randomly
3. Gene flow occurs when individuals leave one population,
join another, and breed. Is like migration
4. Mutation modifies allele frequencies by continually
introducing new alleles. Generate new alleles within a pop.

Changes in allele frequency

what can they tell us? Predictions?
Alleles have frequencies within populations
These frequencies can change from generation to generation
this is the process of evolution
The factors that cause changes in allele frequency are the causes
of evolution (we just went over them)
To study these causes, it is useful to have a null model a
model of no changes
The null is useful because we can test for departures from it,
therefore determining evolution has occurred, and then we can
test for the causes

Hardy-Weinberg Principle
Modern synthesis refers to the era in which
scientists of many disciplines collaborated to
quantify evolution
One product of this era is the model known
as the HardyWeinberg principle
In 1908, G. H. Hardy and Wilhelm Weinberg
Studied how the four evolutionary processes
affect populations
Wanted to know what happened in an entire
population when all of the individualsand
thus all possible genotypesbred
Developed a mathematical model to analyze
the consequences of such matings

Hardy-Weinberg Principle
It is called the Hardy-Weinberg Principle
This principle states that whatever the initial frequencies for two alleles must be, after one generation of random mating, the
frequencies will be:

p2 : 2pq : q2
In a large randomly breeding population, allelic frequencies
and
will remain the same
fromthus
generation to generation
assuming that there is no mutation, gene migration, selection
or genetic drift.
2

p2 + 2pq + q = 1

The Gene Pool Concept

Hardy and Weinberg imagined that all of the gametes
produced in each generation go into a single group called
a gene pool and then combine randomly

Their calculations predict the genotypes of the offspring

that the population would produce, as well as the
frequency of each genotype

Deriving the HardyWeinberg Principle

They started with the simplest situation, a gene with two
alleles, A1 and A2
The frequency of A1 is represented by p and the frequency of
A2 is represented by q. Because there are only two alleles,
pq1
In this situation, three genotypes are possible: A1A1, A1A2, and
A2A2. The model predicts the following genotype frequencies:
The frequency of the A1A1 genotype is p2
The frequency of the A2A2 genotype is q2
The frequency of the A1A2 genotype is 2pq

Deriving the HardyWeinberg Principle

Because all individuals in the new generation must have
one of the three genotypes, the sum of the three genotype
frequencies must equal 1 (100% of the population):
p2 2pq q2 1
This is the HardyWeinberg equation
When allele frequencies are calculated for this new
generation, the frequency of A1 is still p and the frequency
of A2 is still q

p2 : 2pq : q2

Homozygote 1
(AA)

Heterozygote
(Aa)

Homozygote 2
(aa)

Frequencies
Frequencies at
at this
this ratio
ratio are
are said
said to
to be
be in
in Hardy-Weinberg
Hardy-Weinberg
equilibrium
equilibrium
The
The frequencies
frequencies will
will remain
remain as
as such
such unless
unless factors
factors not
not yet
yet
discussed
discussed (such
(such as
as natural
natural selection)
selection) change
change them
them

Genotype and allele frequencies for a

locus with two alleles

Genotype and allele frequencies for a

locus with two alleles

2pq

p2

q2

Genotype and allele frequencies for a locus with two

alleles

Genotype and allele frequencies for a locus with two

alleles

Genotype and allele frequencies for a locus with two

alleles

2pq
p2 q2

p2

2pq

q2

When alleles are transmitted via meiosis and random combination of

gametes, their frequencies do not change over time

Important Assumptions of the

HardyWeinberg Model

1. Allele frequencies: p q 1
2. Genotype frequencies: p2 2pq q2 1

The HardyWeinberg model is based on assumptions

about the behavior of populations and alleles
The assumptions helped to define the four processes of
evolution that can act on a population

Important Assumptions of the HardyWeinberg

Model
The HardyWeinberg principle makes predictions based on
these assumptions:
1.
2.
3.
4.
5.

Random mating
No natural selection
No genetic drift (random allele frequency changes)
No gene flow
No mutation

Under these conditions:

The genotypes A1A1, A1A2, and A2A2 will be in the Hardy
Weinberg proportions p2, 2pq, and q2
No evolution will occur

Important Assumptions of the Hardy

Weinberg Model
Biologists often want to test whether
natural selection is acting on a particular gene
nonrandom mating is occurring
one of the other evolutionary mechanisms is at work
In addressing questions like these, the Hardy Weinberg
principle functions as a null hypothesis
When genotype frequencies do not conform to Hardy
Weinberg proportions, evolution or nonrandom mating is
occurring in that population

Nonrandom mating
Assortative mating individuals choose
to mate with others who are more
phenotypically similar (or less similar) to
themselves than would be expected on a
random basis.
Positive assortative mating will result in
increased number of homozygotes
Negative assortative mating will result in
increased number of heterozygotes.

Inbreeding nonrandom mating based on genetic relatedness.

- Homozygous parents that self-fertilize produce all homozygous
offspring

Inbreeding
In each generation, inbreeding
Increases the frequency of homozygotes
Reduces the frequency of heterozygotes
Inbreeding itself does not cause evolution, because allele frequencies
do not change in the population as a whole
Nonrandom mating changes genotype frequenciesnot allele
frequencies
Nonrandom mating itself is thus not an evolutionary process

How Does Inbreeding Influence

Evolution?
Although inbreeding does not directly cause evolution, it
can speed the rate of evolutionary change
It increases the rate at which natural selection
eliminates deleterious alleles from a population
Inbreeding depression is a decline in average fitness
that takes place when homozygosity increases and
heterozygosity decreases in a population

How Does Inbreeding Influence

Evolution?
Inbreeding depression results from two processes:
1. Many recessive alleles represent loss-of-function
mutations
In heterozygotes, these alleles have little or no effect
Inbreeding increases the frequency of homozygous recessive
individuals

2. Many genesespecially those involved in fighting disease

are under intense selection for heterozygote advantage
If an individual is homozygous at these genes, then fitness
declines

Case Study: Florida Panthers

Percent survivorship of offspring

(a component of fitness)

Figure 26.4

Texas
Florida
panthers
Florida
Florida
panthers
Age (yr)

Among panthers, offspring had decreased fitness when

both parents were from Florida compared to those who
had one parent from Florida and one from Texas

Nonrandom mating
sexual selection
Another form of nonrandom mating is sexual selection.
Sexual selection (Darwin wrote a book on this too!) if
individuals are choosy for certain traits it wont be random

Intrasexual selection directly compete for mating

Intersexual selection mate choice showy characteristics
Sexual selection does lead to changes in allele frequencies
in a population
It is a form of natural selection

Nonrandom mating
sexual selection
Sexual selection violates the assumptions of the Hardy
Weinberg principle
Causes certain alleles to increase or decrease in
frequency
Results in evolutionary change
Other types of natural selection favor traits that help
organisms survive in their environments
Natural selection is the only evolutionary process that
results in adaptation

Four Agents of Evolutionary Change

1. Selection some genotypes do better than others in
their environment
2. Genetic drift the effect of chance on small populations
3. Gene flow movement of alleles between populations
4. Mutation ultimate source of variation

Any of these forces may bring about changes in allele

frequencies. Multiple forces can act at once!

Four Agents of Evolutionary Change

1. Selection some genotypes do better than others in
their environment
2. Genetic drift the effect of chance on small populations
3. Gene flow movement of alleles between populations
4. Mutation ultimate source of variation

Any of these forces may bring about changes in allele

frequencies. Multiple forces can act at once!

Natural Selection

Natural selection occurs when

Heritable variation leads to differential reproductive success
Individuals with certain phenotypes produce more surviving
offspring than individuals with other phenotypes do.

The result is evolution

HardyWeinberg model assumption has been violated

How Does Selection Affect Genetic

Variation?
Three conditions must be met for evolutionary change to occur:
1. Variation must exist among individuals in a population.
Genetic variation - the number and relative frequency of alleles that are
present in a particular population
Lack of variation can make populations less able to respond
successfully to changes in the environment

2. Variation must be genetically inherited

3. This variation must result in different fitness for different
individuals (i.e. because of their phenotype, some individuals
will be more successful and will have more offspring than
others with a different phenotype)

Selection
Selection can cause adaptation to many different
pressures
- Physiological tolerances to environment climate,
chemicals, etc.
- Avoiding detection (as prey or predator)

Snowy owl

Red Underwing moth

Katydid

Ring-necked snake

Zebra

Red Underwing moth

How Does Selection Affect Genetic

Variation
Natural selection occurs in a wide
variety of patterns:
Directional
Stabilizing
Disruptive
Balancing selection (not shown)

Directional selection
Evolution causes
an increase in the
number of
individuals with the
trait at one end of
the curve.
= Directional
selection reduces
the genetic
diversity of
populations

Peccary

Stabilizing selection
Individuals in the center of the
curve and more fit and thus the
curve narrows
= Stabilizing selection
reduces genetic variation in
a trait
Average value of a trait
does not change over
time
Reduces both extremes in
a population

Disruptive selection
Individuals at the ends of the
curve are more fit than the
ones in the middle causing
two curves to form and
possible new species to
form.
= Disruptive selection
increases variation in a trait.
Intermediate phenotypes are
selected against and
extreme phenotypes are
favored

Balancing Selection
Balancing selection maintains variation in a trait
Occurs when no single allele has a distinct advantage
There is a balance among several alleles in terms of
their fitness and frequency

Selection can maintain variation

(balancing selection)
Balancing selection occurs when
Frequency-dependent selection
Negative (rare is good!)
Positive (rare is bad!)
Can be complex
Oscillating selection
Environmental fluctuations
Heterozygote advantage
Neither homozygote is as fit

Sickle cell anemia

-Heterozygote
advantage

Natural Selection Summary

No matter how natural selection

occurs, it increases fitness and leads
to adaptation
Natural selection results in allele
frequencies that deviate from those
predicted by the HardyWeinberg
principle
Selection favors certain alleles
over others

Four Agents of Evolutionary Change

1. Selection some genotypes do better than others in
their environment
2. Genetic drift the effect of chance on small populations
3. Gene flow movement of alleles between populations
4. Mutation ultimate source of variation

Any of these forces may bring about changes in allele

frequencies. Multiple forces can act at once!

Genetic Drift
Genetic drift is any change in allele frequencies in a population due
to chance (sampling error)
It causes allele frequencies to drift up and down randomly over
time
Drift occurs in every population, in every generation
It is especially prevalent in small populations
Genetic drift has a much larger effect on small populations than on
large populations
Over time, one allele will drift to fixation or loss
Given enough time, drift can still be an important factor in large
populations

Genetic Drift
Chance can have a major
impact on small populations
Alleles can be lost easily
even alleles which could be
beneficial!
New mutations could die out or
rise to high frequency
Can potentially negate natural
selection

What Causes Genetic Drift in Natural

Populations?
Genetic drift can be caused by any event or process that
involves sampling error
Two examples are founder effects and bottlenecks

Founder Effects
A founder event occurs when a group starts a new
population in a new area
Especially in small groups, allele frequencies probably differ
from the source population
This change in allele frequencies is called a founder
effect
Founder effects are especially common in the colonization of
isolated habitats
Each time a founder event occurs, a founder effect is likely to
accompany it, changing allele frequencies through genetic
drift

Founder Effect

Red = 0.50, Green = 0.50

Red = 0.20, Green = 0.80

Founder effect is a form of genetic drift

Genetic Bottlenecks
A sudden decrease in population size is called a population
bottleneck
Population bottlenecks are commonly caused by disease
outbreaks and natural catastrophes
Population bottlenecks lead to genetic bottlenecksa
sudden reduction in the number of alleles in a population
Drift occurs during genetic bottlenecks and causes a change
in allele frequencies

Bottleneck Effect

Bottleneck effect is a form of genetic drift

Bottleneck Effect

Same population size, but NOT the same

amount of genetic variation

Bottleneck Effect

Northern elephant seals

Bottleneck: human hunting

Four Agents of Evolutionary Change

1. Selection some genotypes do better than others in
their environment
2. Genetic drift the effect of chance on small populations
3. Gene flow movement of alleles between populations
4. Mutation ultimate source of variation

Any of these forces may bring about changes in allele

frequencies. Multiple forces can act at once!

Gene Flow

Gene Flow - Movement of alleles from one population to another

Could be due to individuals physically moving - or their gametes

Can change allele frequencies

Greatly affected by mobility of the organism, barriers to dispersal

Without gene flow populations could end up becoming genetically

differentiated to the point of becoming different species

Measuring Gene Flow between

Populations
The decline of gene flow between isolated wild
populations is well documented
Isolating events include habitat fragmentation
Gene flow between wild and captive populations is also
well documented

Measuring Gene Flow between

Populations
Biologists studied steelhead trout in the Hood River
of Oregon
Some of the trout are wild
Some of the trout were raised in a hatchery and released

DNA tests were conducted to study the reproductive fitness of

Individuals with two wild parents
Individuals with one wild and one captive-bred parent
Individuals with two captive-bred parents
(a) Wild steelhead trout populations are declining

Measuring Gene Flow between

Populations
On average, compared to the fitness of fish with two wild
parents:
Fish with one captive-bred parent have 16% lower fitness
Fish with two captive-bred parents have 38% lower fitness

Augmenting wild populations with captive-bred fish may

decrease, rather than increase, population size

Gene Flow Is Random with Respect to

Fitness
Gene flow does not always reduce fitness
It can replenish alleles in a population that has lost
alleles due to genetic drift
This should increase genetic diversity
Gene flow can increase or decrease fitness depending on
the situation
However, movement of alleles between populations
always tends to reduce their genetic differences

Mutation
Most evolutionary mechanisms reduce genetic diversity over
time
However, mutation restores genetic diversity by creating
new alleles
There are several ways in which a mutation can occur:
Point mutations
Chromosome-level mutations
Lateral gene transfer

Mutation
Mutation is an evolutionary process that increases genetic diversity in
populations
Mutation is random with respect to the fitness of the individual with
the affected allele
In other words, mutation does not occur in a way that tends to
increase or decrease fitnessmutation just happens
Most mutations result in deleterious alleles, alleles that lower fitness
These alleles tend to be eliminated by purifying selection
Rarely, mutation produces a beneficial allele
These alleles should increase in frequency in a population due to
natural selection

Mutation as an Evolutionary Mechanism

In principle, mutation can change the frequencies of alleles
over time
However, mutation alone is rarely enough to make it an
important factor in changing allele frequencies
Mutation can be a significant evolutionary force in bacteria
and archaea, which have short generation times
As an evolutionary mechanism, mutation is relatively slow
compared with natural selection, genetic drift, and gene flow
When combined with other evolutionary mechanisms,
however, mutation can have a large effect on evolution

Experimental Studies of Mutation

The bacterium Escherichia coli has been used as a model to
study how mutation affects evolution
In one experiment, Richard Lenski and colleagues set up
populations of E. coli and followed them for 10,000
generations
E. coli is asexual, so mutation is its only source of genetic
variation
No gene flow occurred
Both selection and genetic drift operated in each
population

Experimental Evolution
- Bacteria
Can Bacterial Evolve?
Yes
Mutation
Bacteria make excellent models to study evolution
Easy to grow in a laboratory
Short generation time
Can be frozen a different points, then regrown
Re-evolve
Evolution can be observed directly

Experimental Evolution
- Bacteria
E. coli long-term evolution experiment
Richard Lenski, 24 Feb 1988
12 genetically identical populations
Early years
All populations evolved larger cell volumes
Lower maximum population densities
Became specialized for living on glucose
4 population developed defects in DNA repair
Increased mutation rate

Experimental Evolution
- Bacteria
E. coli long-term evolution experiment
2008
1 population
Ability to grow on citrate
Visible in generation 33,127
Citrate using variant evolved at some point between
generation 31,000 and 31,500
Re-evolve from these generations
Not able to re-evolve before 20,000

Experimental Evolution
- Bacteria

The researchers found that the relative fitness of the populations

increased over time in jumps
They suggested that this pattern resulted from novel mutations
arising and conferring a fitness benefit
Genetic drift probably had a small effect because the
populations were large

Studies of Mutation in Natural

Populations
Pea aphids are small insects that feed on plant sap
There are two phenotypesred and greenin wild populations
Color is determined by carotenoid
pigments
Pigment does not come from plant sap
Aphids are the first animals ever discovered
to be able to generate their own carotenoids
The genes that code for the metabolic
pathway came from mutations
Lateral gene transfer from a fungal symbiont
Additional mutations occurred after the gene transfer

Take-Home Messages
Mutation plays an important role in evolution:
1. Mutation is the ultimate source of genetic variation
Crossing over and independent assortment shuffle existing
alleles into new combinations
Only mutation creates new alleles
2. Without mutation, evolution would eventually stop
3. Mutation alone is usually inconsequential in changing allele
frequencies at a particular gene
Each of the four evolutionary forces has different consequences for
allele frequencies
All four evolutionary mechanisms violate HardyWeinberg predictions
The ultimate result of these mechanisms is biological diversity

Summary
Summary Table 26.4